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Higher plants exhibit a variety of different life histories. Annual plants live for less than a year and after flowering produce seeds and senesce. By contrast perennials live for many years, dividing their life cycle into episodes of vegetative growth and flowering. Environmental cues control key check points in both life histories. Genes controlling responses to these cues exhibit natural genetic variation that has been studied most in short-lived annuals. We characterize natural genetic variation conferring differences in the perennial life cycle of  Arabis alpina . Previously the accession Pajares was shown to flower after prolonged exposure to cold (vernalization) and only for a limited period before returning to vegetative growth. We describe five accessions of  A. alpina  that do not require vernalization to flower and flower continuously. Genetic complementation showed that these accessions carry mutant alleles at  PERPETUAL FLOWERING 1  ( PEP1 ), which encodes a MADS box transcription factor orthologous to FLOWERING LOCUS C in the annual  Arabidopsis thaliana . Each accession carries a different mutation at  PEP1 , suggesting that such variation has arisen independently many times. Characterization of these alleles demonstrated that in most accessions, including Pajares, the  PEP1  locus contains a tandem arrangement of a full length and a partial  PEP1  copy, which give rise to two full-length transcripts that are differentially expressed. This complexity contrasts with the single gene present in  A. thaliana  and might contribute to the more complex expression pattern of  PEP1  that is associated with the perennial life-cycle. Our work demonstrates that natural accessions of  A. alpina  exhibit distinct life histories conferred by differences in PEP1 activity, and that continuous flowering forms have arisen multiple times by inactivation of the floral repressor  PEP1 . Similar phenotypic variation is found in other herbaceous perennial species, and our results provide a paradigm for how characteristic perennial phenotypes might arise.

PEP1 of Arabis alpina Is Encoded by Two Overlapping Genes That Contribute to Natural Genetic Variation in Perennial Flowering